12 4 1920 the journal of data 8 July 2016 Check List Lists of Species Check List 12(4): 1920, 8 July 2016 doi: http://dx.doi.org/10.15560/12.4.1920 ISSN 1809-127X © 2016 Check List and Authors

Arachnids from Araripe Plateau, Ceará, Brazil

Raul Azevedo1*, Eridiane da S. Moura1, Adreany S. Lopes2, Leonardo S. Carvalho3, 4, Sidclay C. Dias5 and Antonio D. Brescovit 6

1 Universidade Federal do Cariri, Laboratório de Entomologia. Rua Vereador Maciel Lopes, s/n, São José, CEP 63133-630, Crato, CE, Brazil 2 Quadra 15, Bloco 03, apto. 302, Morada Nova, CEP 264023-210, Teresina, PI, Brazil 3 Universidade Federal do Piauí, Campus Amílcar Ferreira Sobral – CAFS, BR 343, km 3.5, Bairro Meladão, CEP 64800-000, Floriano, PI, Brazil 4 Programa de Pós-Graduação em Zoologia, Universidade Federal de Minas Gerais, Belo Horizonte, MG, Brazil 5 Rua Arício Guimarães Fortes, Ed. Jardim da Atalaia, 826, apto. 401, CEP 49037-060, Atalaia, Aracaju, SE, Brazil 6 Laboratório Especial de Coleções Zoológicas, Instituto Butantan. Av. Vital Brasil, 1500 05503-900 São Paulo, SP, Brazil * Corresponding author. E-mail: [email protected]

Abstract: The Araripe Plateau is situated in the states harvestmen (De Souza et al. 2014) species recorded for of Ceará, Piauí and Pernambuco, northeastern Brazil, the Caatinga biome. These numbers represent about in the Caatinga biome, semiarid domain. This region 8.5%, 21% and 5%, respectively, of the , scor- of Brazil is characterized by high temperatures and pions and Laniatores harvestmen species recorded in low rainfall. Nevertheless, due to its high elevation Brazil (Carvalho et al. 2014; Porto et al. 2014; De Souza and large amount of orographic rainfall, the Araripe et al. 2014). The potential explanations for these num- Plateau is an exception as a seasonal semi-deciduous bers are different for each group. Considering that the forest with high humidity in the semiarid domain and Caatinga occupies about 10% of Brazilian territory, the the Caatinga biome. Herein a species list of Araripe number of species might represent a clear effect Plateau is presented from the compilation of sampling deficiency, as it is lower than expected by of data from in situ sampling from January to May chance (Carvalho et al. 2014). The scorpion species rich- 2013, from the literature, and from Brazilian ness in the Caatinga is higher than expected by chance. collections data. A total of 68 species were recorded for However, considering that the world’s highest scorpion the area, of which 53 were spiders, eight harvestmen species richness (4–13 sympatric species at one site) and seven scorpions. The species richness recorded here occurs in drier areas (Polis 1990), more scorpion species is considered significant regarding the known diversity would be expected to be found in the Caatinga. On the of Arachnida in the semiarid region and contributes to other hand, for harvestmen, the low species richness expanding our knowledge of the area. in the Caatinga biome was expected, as many species are ineffective at preventing water loss (Santos 2007a), Key words: Arachnida; Araneae; Scorpiones; thus resulting in higher species richness in humid areas Opiliones; Caatinga throughout Brazil, such as the Atlantic Forest (Pinto-da- Rocha et al. 2005). In the present paper, we present a list of the spiders, INTRODUCTION scorpions and harvestmen from the Araripe Plateau, The Caatinga biome is one of the least studied northeastern Brazil. The arachnid fauna of the area has Neotropical biomes (Leal et al. 2005; Santos et al. been the focus of only a few published papers, mainly 2011), with current data suggesting that about 41% consisting of the description of fossil (i.e., Campos of the area has never been surveyed by scientists and 1986; De Carvalho and Lourenço 2001; Selden et al. 80% has been poorly sampled (Tabarelli and Vicente 2006) or living taxa (i.e., Bonaldo and Brescovit 1998; 2004). This situation is even worse for taxa such as Corronca 1998; Rheims and Brescovit 2004; Lourenço invertebrates (Lewinson et al. 2005). Among arachnids, 2010, 2014; Ruiz 2013; Huber et al. 2014). None of recent estimates indicate that there are no records of these publications directly dealt with the Araripe spiders (Carvalho et al. 2014) or scorpions (Porto et al. Plateau arachnid fauna, but focused on taxonomic 2014) for about 70% of the Caatinga area. issues rather than providing species lists. The present Even considering such heterogeneous sampling, paper is the first to compile published records and new presently there are 271 spiders (Carvalho et al. 2014), records gathered from field work into a species list for 28 scorpions (Porto et al. 2014) and 28 Laniatores the area.

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Figure 1. Map of the sampled localities (black dots) and its location in the Araripe National Forest (in green) and the Araripe Plateau (in grey). Municipali- ties below the thick line are located in the State of Pernambuco, and those above are located in the State of Ceará.

MATERIALS AND METHODS (with and without UV lamps) and litter sampling Study area (1 m2 of litter sifted and hand sorted in each sample). The Araripe Plateau is located in northeastern Brazil, All standardized samplings followed usual arachnid in the states of Ceará, Piauí and Pernambuco. It includes sampling methods, as described in Carvalho (2015). the Araripe National Forest (Floresta Nacional do Ara- The species were identified using keys for spiders (Levi ripe, 07°20ʹ42ʺ S, 039°25ʹ6ʺ W). The region is within 1968, 1991, 1997, 2004; Corronca 1998; Bonaldo 2000; the Caatinga biome, a warm (mean annual temperature Brescovit and Rheims 2000; Rheims and Brescovit 2004; 26°C − 28°C; Nimer 1972) semiarid biome. Despite the Santos 2007b; Abrahim et al. 2012), scorpions (Lourenço dry and warm environment, the high elevation of the 2002; De Souza et al 2009) and harvestmen (Pinto-da- region (more than 500 m above sea level), contributes to Rocha 1997) and by comparison with identified material the production of approximately 1,200 mm of orograph- deposited in institutional collections. ic rainfall (Andrade-Lima 1982; Prado 2003; Leal et al. Collected specimens were deposited in the following 2005). This allows the formation of forested enclaves collections: Museu Paraense Emílio Goeldi, Belém, Pará on the windward slopes of hills, known as “brejo-de- (MPEG; curator A.B. Bonaldo); Instituto Butantan, altitude” (Prado 2003; Leal et al. 2005). São Paulo, São Paulo (IBSP; curator A.D. Brescovit); Universidade Federal de Minas Gerais, Belo Horizonte, Data collection Since 2010, occasional arachnid surveys have been Minas Gerais (UFMG; curator A.J. Santos); Coleção carried out by various researchers in several localities, de História Natural da Universidade Federal do Piauí both inside and outside the Floresta Nacional do Araripe, (CHNUFPI; curator E.F.B. Lima); Universidade Federal such as the municipalities of Crato, Juazeiro do Norte da Paraíba, João Pessoa, Paraíba (UFPB; curator M.B. and Barbalha, state of Ceará, (Figure 1). The sampling da Silva) and Museu Nacional do Rio de Janeiro, Rio de points map (Figure 1) was constructed using ArcGis 10.3 Janeiro, Rio de Janeiro (MNRJ; curator A.B. Kury). software (ESRI 2015). The species list herein also includes information The arachnids were sampled using three techniques: gathered from previous publications on arachnids of pit-fall traps for invertebrates (500 ml plastic buckets the Araripe plateau: Bonaldo and Brescovit (1998), buried at ground level, with 70% ethanol as preservative, Corronca (1998), Lourenço (2010, 2014), Lourenço et placed at least 5 m apart and kept open for at least five al. (2013), Maury (1982), Rheims and Brescovit (2004), consecutive days), diurnal and nocturnal hand searches Ruiz (2013) and Huber et al. (2014).

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RESULTS The compiled list includes 68 arachnid species, C. L. Koch, 1841 (), Neodrassex Ott, 2012 represented by 53 spider, seven scorpions and eight (Gnaphosidae), Otiothops MacLeay, 1839 (Palpiman­ harvestmen species (Table 1). Thirty-five species idae), Ariadna Audouin, 1826 (Se­gestri­idae) and Tenedos were identified to species level and six un­described O. Pickard-Cambridge, 1897 (Zodariidae). The spider species were recognized, belonging to the genera family Corinnidae and scorpion family Buthidae pre- Orthonops­ Chamberlin, 1924 (), Corin­na sented the highest species richness.

Table 1. Spider, scorpion and harvestman species list from the Araripe Plateau, Ceará, Brazil, with the number of specimens collected and source of information gathered.

Specimens from museums or Taxa Total Species collected in this study cited in the literature ARANEAE Araneidae Araneus venatrix (C. L. Koch, 1838) 1 MPEG 024478 Aculepeira cf. travassosi (Soares & Camargo, 1948) 1 CHNUFPI 1203 Araneus guttatus (Keyserling, 1865) 2 CHNUFPI 1205, 1225 Argiope argentata (Fabricius, 1775) 1 CHNUFPI 1210 Kapogea cyrtophoroides (F. O. Pickard-Cambridge, 1 CHNUFPI 1219 1904) Caponiidae Orthonops sp. n. 1 2 MPEG 24475, 024476 Corinnidae gr. aenea sp. n. 2 3 CHNUFPI 0102, 0127, 0136 Corinna gr. capito sp. 1 1 CHNUFPI 0119 Falconina gracilis (Keyserling, 1891) 2 MPEG 024479, 024480 Abapeba gr. abalosi (Mello-Leitão, 1942) 2 MPEG 024437, 024438 Attacobius verhaaghi Bonaldo & Brescovit, 1998 1 Bonaldo and Brescovit (1998) Corinna sp. 1 3 MPEG 024447, 024448, 024449, sp. 1 4 MPEG 024433, CHNUFPI 0113 sp. 1 1 MPEG 024435 Castianeira sp. 2 1 MPEG 024432 Corinninae sp. 1 1 CHNUFPI 0092 Castianeirinae sp. 1 1 MPEG 024434 Ctenidae Ctenus sp. 1 1 CHNUFPI 1217, 1224 Cyrtaucheniidae Fufius sp. 1 1 CHNUFPI 1218 Dictyna sp. 1 1 CHNUFPI 1211 Deinopidae Deinopis sp. 1 2 CHNUFPI 1081 Gnaphosidae Gnaphosidae sp. 1 1 MPEG 024472 Neodrassex sp. n. 1 1 MPEG 024470 Hahniidae Hahniidae sp. 1 1 MPEG 024450 Hersillidae Ypypuera crucifera (Vellard,1924) 2 CHNUFPI 0435 Rheims & Brescovit (2004) Linyphiidae Agyneta sp. 1 4 CHNUFPI 1215, 1220,1221, 1223 Gamasomorpha lutzi (Petrunkevitch, 1929) 2 MPEG 024445, 024446 Neoxyphinus termitophilus (Bristowe, 1938) 3 MPEG 024467, 024468, 024469 Nephilidae Nephila clavipes (Linnaeus, 1767) 1 CHNUFPI 1208 Palpimanidae Otiothops sp. n. 1 2 MPEG 024474, 024471, 024477 Pisauridae Architis tenuis Simon, 1898 7 CHNUFPI 1200, 1214, 1222 sp. 1 1 CHNUFPI 0103, 0141 Orthobula sp. 2 1 CHNUFPI 0079, 0113, 0091 Continued

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Table 1. Continued.

Specimens from museums or Taxa Total Species collected in this study cited in the literature Pholcidae Micropholcus crato Huber, Carvalho & Benjamin, 2014 4 Huber et al. (2014) Salticidae Soesilarishius macrochelis Ruiz, 2013 1 MPEG 20278 Soesilarishius cearensis Ruiz, 2013 1 MPEG 20293 Scytodidae Scytodes eleonorae Rheims & Brescovit, 2001 3 CHNUFPI 0468, 0471, 0472 Segestriidae Ariadna sp. n. 1 2 CHNUFPI 0247, 0248 Selenops maranhensis Mello-Leitão, 1918 2 Corronca (1998) Senoculidae Senoculus sp. 1 1 CHNUFPI 0424 Sicariidae Loxosceles amazonica Gertsch, 1967 1 UFMG 12064 Sicarius cariri Magalhães, Brescovit & Santos, 2013 4 CHNUFPI 0071 IBSP 9142, UFMG 11690, 11691 Tetragnathidae Chrysometa sp. 1 1 CHNUFPI 1201 Leucauge sp. 1 1 CHNUFPI 1204 Leucauge sp. 2 1 CHNUFPI 1212 Theridiidae Chrosiothes sp. 1 2 MPEG 024442, 024443 cf. Theridion sp. 1 1 CHNUFPI 1213 Theridiidae sp. 1 2 MPEG 024441 Theridiidae sp. 2 1 MPEG 024444 Thomisidae Bucranium taurifrons (O. P.-Cambridge, 1881) 1 MPEG 024451 Zodariidae Leprolochus mucuge Lise, 1994 42 MPEG 024452, 024453, 024454, 024455, 024456, 024457, 024458, 024459, 024460, 024461, 024462, 024463, 024464, 024465, 024466 Leprolochus oeiras Lise, 1994 5 MPEG 024439, 024440 Tenedos sp. n. 1 1 MPEG 024473, 024473 SCORPIONES Bothriuridae Bothriurus rochai Mello-Leitão, 1932 1 Maury (1982) Chactidae Hadrurochactas araripe Lourenço, 2010 1 Lourenço (2010) Buthidae Ananteris franckei Lourenço, 1982 11 CHNUFPI 1206, 1207, 1227 Lourenço et al. (2013)

Tityus martinpaechi Lourenço, 2001 1 UFMG 12534 Tityus stigmurus (Thorell, 1877) 8 CHNUFPI 0902, 0908, 0911, 0917, 0921, 1216,1226 UFMG 12533, IBSP 3841 Rhopalurus agamemnon (C.L. Koch, 1839) 1 IBSP 816 Rhopalurus brejo Lourenço, 2014 1 Lourenço (2014) OPILIONES Escadabiidae Escadabiidae sp. 1 1 MNRJ 02121 Cosmetidae Eupoecilaema megaypsilon (Piza, 1938) 3 UFPB 251 Gonyleptidae Pseudopucrolia discrepans (Roewer, 1943) 2 MNRJ 04683, UFPB 255 Parapachyloides uncinatus (Søerensen, 1879) 3 MNRJ 17474, UFPB 253 Sclerosomatidae Munequita sp. 1 2 UFPB 254 Stygnidae Stygnus polyacanthus (Mello-Leitão, 1923) 1 UFPB 252 Protimesius sp. 1 1 MNRJ 05272 Zalmoxidae Zalmoxidae sp. 1 1 MNRJ 02123

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DISCUSSION Selenops maranhensis, B. rochai and R. agamemnom are Attacobius verhaaghi Bonaldo & Brescovit, 1998 widespread species (Corronca 1998; Lourenço 2002) (Corinnidae), Selenops maranhensis Mello-Leitão, 1918 and their absence from our sampling is attributed to (Selenopidae), Bothriurus rochai Mello-Leitão, 1932 sampling bias. Conversely, A. verhaaghi is rare in spider (Bothriuridae), Hadrurochactas araripe Lourenço, 2010 samplings, because most species of the Attacobiini (a (Chactidae), and Rhopalurus brejo Lourenço, 2014 tribe of Corinnidae) are ant-clinging spiders and are (Buthidae) have been reported from the Araripe Plateau expected to live near ant nests, where spider samplings only from literature records, respectively by Bonaldo were rarely performed. The buthid scorpions R. brejo and Brescovit (1998), Corronca (1998), Maury (1982), and H. araripe are both recorded from Araripe based on and Lourenço (2010, 2014). Rhopalurus agamemnon specimens whose precise type localities and preferable (C.L. Koch, 1839) is recorded for the Araripe Plateau microhabitat are unknown (Lourenço 2010, 2014). by a single specimen in the IBSP collection (Table 1). Among the arachnids collected, 22 species were

Figure 2. Representatives of the spiders from the Araripe Plateau, Ceará, Brazil. A: Ctenidae, Ctenus sp.1, female carrying egg-sac; B: Cyrtaucheniidae, Fufius sp.1; C: Deinopidae, Deinopis sp.1; D: Nephilidae, Nephila clavipes, female with kleptoparasitic flies;E: Pisauridae, Architis tenuis; F: Sicariidae, Loxosceles amazonica. Photos: L.S. Carvalho.

Check List | www.biotaxa.org/cl 5 Volume 12 | Number 4 | Article 1920 Azevedo et al. | Arachnids from Floresta Nacional do Araripe identified, most of them widespread in the region (e.g., crato Huber, Carvalho & Benjamin, 2014, Soesilarishius Ypypuera crucifera (Vellard, 1924), Neoxyphinus termi- macrochelis Ruiz, 2013 and Soesilarishius cearensis Ruiz, tophilus (Bristowe, 1938), Architis tenuis Simon, 1898, 2013 are known only from the Araripe Plateau (Lou- Tityus stigmurus (Thorell, 1877); Rheims and Brescovit, renço 2002, 2004; Ruiz 2013; Huber et al. 2014; Porto 2004; Santos 2007b; De Souza et al. 2009; Abrahim et al. 2014). These last four species must be considered et al. 2012). Some of the collected species are com- endemic to the Araripe rainforest, as no further record monly observed in synanthropic environments, such as of them outside that area are known (Giupponi et al. Scytodes eleonorae Rheims & Brescovit, 2001, Loxosceles 2009, Lourenço 2012, 2013; Ruiz 2013; Huber et al. amazonica Gertsch, 1967 (Figure 2F) and Argiope argen- 2014). tata (Fabricius, 1775) (Lucas et al. 1986; Rheims and The Araripe Plateau constitutes a natural forested Brescovit 2001; Levi 2004). On the other hand, Anan- enclave (Prado 2003; Leal et al. 2005), also known as teris franckei Lourenço, 1982 (Figure 3B), Micropholcus “brejo-de-altitude” (Andrade-Lima 1982). The fauna

Figure 3. Representatives of the scorpions and harvestmen from the Araripe Plateau, Ceará, Brazil. A: Bothriuridae, Bothriurus rochai; B: Buthidae, Anan- teris franckei; C: Buthidae, Tityus martinpaechi; D: Buthidae, Tityus stigmurus; E: Cosmetidae, Eupoecilema megaypsilon; F: Gonyleptidae, Parapachyloides uncinatus. Photos: L.S. Carvalho.

Check List | www.biotaxa.org/cl 6 Volume 12 | Number 4 | Article 1920 Azevedo et al. | Arachnids from Floresta Nacional do Araripe and flora of these areas are different from the typical authors would like to thank two anonymous referees for Caatinga environment and usually exhibit a trend valuable comments in a previous version of this paper. toward high diversity (Werneck 2011). These warmer This work is part of the Programa de Pesquisas em environmental conditions are also more suitable for a Biodiversidade do Semiárido, funded by the Conselho myriad of arachnid species which are absent from the Nacional de Desenvolvimento Científico e Tecnológico arid environment surrounding the forested area, such as (CNPq #457471/2012-3). Funding for this research has harvestmen (Curtis and Machado 2007), which prefer been provided by FAPESP grants 2011/50689-0 and more humid environments (Santos 2007a; De Souza et CNPq 301776/2004-0 to ADB. al. 2014). The harvestmen fauna of the Araripe Plateau is LITERATURE CITED composed of eight species, a species richness that falls Abrahim, N., A.B. Bonaldo, A.D. Brescovit, C.A. Rheims, A.J. within the expected number for Brazilian open biomes Santos and R. Ott. 2012. A revision of the Neotropical goblin spider Neoxyphinus Birabén 1953 (Araneae, Oonopidae). (Bragagnolo and Pinto-da-Rocha 2003; Pinto-da-Rocha American Museum Novitates 3743: 1–75. http://hdl.handle. et al. 2005). However, the harvestmen community net/2246/6172 is composed of a mixture of taxa typical of different Andrade-Lima, D. de. 1982. Present-day forest refuges in North- environments. Pseudopucrolia discrepans (Roewer, 1943) eastern Brazil; pp. 245–251, in: G.T. Prance (ed.). Biological is usually found in forested enclaves in northeastern diversification in the tropics. New York: Columbia University Press. Brazil and in the Atlantic Forest in Paraíba and Bonaldo, A.B. 2000. Taxonomia da Subfamília Corinninae (Araneae, Parnambuco (Mendes 2011). Parapachyloides uncinatus Corinnidae) nas Regiões Neotropical e Neártica. Iheringia, (Soerensen, 1879) (Figure 3) is widely distributed Série Zoologia, Porto Alegre, 89: 3–148. doi: 10.1590/S0073- throughout the Cerrado and Caatinga biomes (Kury et al. 47212000000200001 2010), mostly associated with more humid areas (pers. Bonaldo, A.B. and A.D. Brescovit. 1998. On Ecitocobius, a new genus obs.). Conversely, Stygnus polyacanthus (Mello-Leitão, from Central Amazonia with comments on the tribe Attacobiini (Arachnida, Araneae, Corinnidae, Corinninae). Spixiana 21(2): 1923) is widely distributed in northeastern Brazil, often 165–172. http://biodiversitylibrary.org/page/28199405 occurring in drier areas, where it can be found under Bragagnolo, C. and R. Pinto-da-Rocha. 2003. Os Opiliões; pp. termite nests or fallen trunks (Pinto-da-Rocha 1997; 526–536, in: M.I.M.S. Lopes, M. Kirizawa and M.M. da R.F. Pinto-da-Rocha and Carvalho 2009). Melo (eds.). Patrimônio da Reserva Biológica do Alto da Serra de This work constitutes the first species list of arachnids Paranapiacaba: a antiga Estação Biológica do Alto da Serra. São Paulo: Instituto de Botânica. for Araripe Plateau. Although the number of identified Brescovit, A.D. and C.A. Rheims. 2000. On the synanthropic species or the total species richness is not especially species of the genus Scytodes Latreille (Araneae, Scytodidae) of high, the records herein presented are important as Brazil, with synonymies and records of these species in other they fill a gap in the known geographical distribution Neotropical countries. Bulletin of the British Arachnological of many arachnid species in the Caatinga biome. Society 11: 320–330. Campos, D.R.B. 1986. Primeiro registro fóssil de Scorpionoidea na These records are important data for proposing and Chapada do Araripe (Cretáceo do Brasil). Anais da Academia delimiting areas of endemism (see Oliveira et al. 2015), Brasileira de Ciências 58(1):135–137. one of the most important tools for selecting priority Carvalho, L.S. 2015. Aracnídeos: quem são, por que estudá-los e areas for conservation. This list is also a first step for como coletá-los?; pp.103–140, in: M.S.C.S. Lima; L.S. Carvalho further ecological and taxonomical investigation on the and F. Prezoto (orgs.). Métodos em Ecologia e Comportamento . Teresina, EDUFPI. 132 pp. arachnids of the region. The huge Wallacean shortfall Carvalho, L.S., A.D. Brescovit, A.J. Santos, U. Oliveira and J.P.L. related to invertebrates from the Caatinga biome Guadanucci. 2014. Aranhas da Caatinga; pp. 15–33, in: F. Bravo prevents the proposition of such wide studies, while with and A. Calor (eds.). Artrópodes do semi-árido: biodiversidade e plants, for example, Caatinga ecoregions were proposed conservação. Feira de Santana: Printmídia. more than a decade ago (Velloso et al. 2002). Thus, it is Corronca, J.A. 1998. The South American spiders of the genus Selenops (Araneae, Selenopidae) with a description of three new expected that with the increase in number of published species. Studies on Neotropical Fauna and Environment 33: papers on geographical distribution, species 124–148. doi: 10.1076/snfe.33.2.124.2153 inventories and new records from the Caatinga, more Curtis, D.J. and G. Machado. 2007. Ecology; pp. 280–308 in: R. ecological, taxonomic and biogeographic studies can be Pinto-da-Rocha, G. Machado and G. Giribet. (eds.). Harvestmen: conducted. the biology of Opiliones. Cambridge: Harvard University Press. De Carvalho, M. da G.P. and W.R. Lourenço. 2001. A new family of fossil scorpions from the Early of Brazil. Earth and ACKNOWLEDGEMENTS Planetary Sciences 332: 711–716. doi: 10.1016/S1251-8050(00) We thank Alexandre B. Bonaldo and Adriano B. Kury 01501-9 for providing data from MPEG and MNRJ arachnids De Souza, A.M., M.B. da Silva, L.S. Carvalho and U. Oliveira. 2014. Opiliões laniatores do semiárido; pp. 47–57, in: Bravo, F. and from FLONA Araripe. We also thank Jimmy Cabra A. Calor (eds.). Artrópodes do semi-árido: biodiversidade e (USP), Rafael Yuji Lemos (IBSP) and Bárbara T. Faleiro conservação. Printmídia: Feira de Santana. (UFMG) for their help in species identification. The De Souza, C.A.R., D.M. Candido, S.M. Lucas and A.D. Brescovit.

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Veloso, A.G., E.V.S.B Sampaio, A.M. Giulietti, M.R.V. Barbosa, Author contributions: RA collected, sorted and analyzed data, and A.A.J.F. Castro, L.P.de Queiroz, A. Fernandes, D.C. Oren, L.A. wrote the text; ESM and ASL collected and sorted data; LSC collected Cestaro, A.J.E. de Carvalho, F.G.C Parein, F.B.R. da Silva, E.E. data, analyzed and wrote the text; SCD and ADB wrote the text. de Miranda, S. Keel and R.S. Gondim. 2002. Ecorregiões pro- postas para o bioma caatinga. A.L. Veloso, E.V.S.B. Sampaio, F.G.C. Pareyn. (eds.). Recife: Associação de plantas de Nordeste. Instituto de Conservação Ambiental The Nature Conservancy do Brasil. 76 pp. Werneck, F.P. 2011. The diversification of eastern South American Received: 23 October 2015 open vegetation biomes: Historical biogeography and perspec- Accepted: 27 April 2016 tives. Quaternary Science Reviews 30: 1630–1648. Academic editor: Gustavo Silva de Miranda

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